Journal of the Marine Biological Association of the United Kingdom, 2009, 89(1), 179–201. #2008 Marine Biological Association of the United Kingdom doi:10.1017/S0025315408002476 Printed in the United Kingdom

REVIEW

Potential impacts of climate change and greenhouse gas emissions on Mediterranean marine ecosystems and cetaceans d.d. gambaiani1, p. mayol1, s.j. isaac2 and m.p. simmonds3 1Souffleurs d’Ecume, Mairie de la Celle, 83170 La Celle, France, 2Halcrow, Burderop Park, Swindon, Wiltshire, SN4 0QD, 3The Whale and Dolphin Conservation Society, Brookfield House, Chippenham, Wiltshire, SN15 1LJ, UK

The combustion of fossil fuels and the resultant impacts on climate may now represent one of the largest environmental threats. In the Mediterranean Sea, changes in bio-chemical and physical seawater properties resulting from global warming are likely to alter marine biodiversity and productivity, trigger trophic web mismatches and encourage diseases, toxic algal bloom and propagation of thermophilic species. This review highlights the current and potential threats of climate change to the Mediterranean marine ecosystems, including cetaceans, and stresses the emergent necessity for more integrated regulations and policies for the protection of marine biodiversity. For instance, in the Mediterranean Sea, the dis- tribution and abundance of the small euphausid species Meganyctiphanes norvegica is correlated with specific hydrobiolo- gical parameters including seawater temperature, salinity and current patterns. Situated at the northern limit of its ecological tolerance, this species, which constitutes the only known food supply of the fin whales (Balaenoptera physalus) in this region, might be affected by climate change-induced alteration of ocean circulation.

Keywords: potential impacts, climate change, greenhouse gas emissions, Mediterranean, marine ecosystems, cetaceans

Submitted 31 July 2007; accepted 3 June 2008; first published online 23 September 2008

INTRODUCTION Intergovernmental on Climate Change (IPCC) shows that, according to different scenarios, the average warming of the Until recently, scientists have hesitated to associate the global planet will be between 1.8 and 48C by 2100 (IPCC, 2007b). warming phenomenon with anthropogenic greenhouse gas Such an increase of temperature in a very short time period emissions. However, a strong correlation has been demon- represents a real ‘climatic shock’ for which no solution has strated along with other supporting evidence and it is now yet been considered by our society (Jancovici, 2005). generally accepted that greenhouse gas emissions are respon- Positive feedback processes would tend to accelerate the rate sible for global warming (e.g. Tett et al., 2000; Smith et al., of global warming (Woodwell & Mackenzie, 1995). The amount 2005; IPCC, 2007a). We know that our carbon dioxide emis- of methane sequestered as methane hydrates in the oceanic sedi- sions, which represent 7 gigatons (Gt) of carbon per year ments of continental plates, as well as in permafrost regions, (Bopp et al., 2002), are responsible for disastrous ecological could be 3000 times larger than the amount of methane disorders. Each year, 2 Gt of carbon is absorbed by the present in the atmosphere (Blunier, 2000). With an increase of oceans through physical and biological processes but, this water temperatures, methane hydrates could become unstable ‘carbon pump’ could be saturated by 2020–2030 (Orr et al., and release vast quantities of methane into the atmosphere. 2001). A significant reduction in the efficiency of the Given that methane is 20 times more efficient than carbon Southern Ocean sink of CO2 has recently been observed dioxide at trapping infrared radiation, this could have cata- (Le Que´re´ et al., 2007). Studies indicate that climate/carbon strophic climatic consequences. Such an event, known as a cycle feedbacks could be responsible for an additional increase ‘methane burp’, occurred about 200 million years ago. By of the partial pressure of CO2 in the atmosphere of 10% increasing the atmospheric temperature by 4 to 8 degrees, and to 50% (Cox et al., 2000). Working Group 1 of the reducing the amount of oxygen, this phenomenon may have been responsible for the extinction of 80% of the oceanic

Corresponding author: fauna at that time (Hesselbo et al., 2000; Kennett et al., 2000). D.D. Gambaiani Atmospheric carbon dioxide is also known to be absorbed by Email: delphine.gambaiani@souffleursdecume.com the oceans, leading to ocean acidification. This process is already

179 180 d.d. gambaiani et al.

affecting surface waters (Royal Society, 2005) and future Furthermore, from 1900 to 2000, the Mediterranean experi- increases in atmospheric carbon dioxide via the combustion of enced a 10% decrease in summer rainfall (IPCC, 2001a). These fossil fuels are expected to profoundly affect ocean chemistry trends are expected to persist and intensify over time, and and marine life (Caldeira & Wickett, 2003; Feely et al.,2004; extreme meteorological events such as droughts (e.g. in the Royal Society, 2005; Orr et al., 2005; Bass et al.,2006). summers of 1995 and 2003) and flooding will become more fre- Global warming is already affecting marine ecosystems quent in the future (e.g. Parry, 2000; IPCC, 2001b; Li, 2003; (CBD, 2003; Parmesan & Yohe, 2003). According to Thomas EEA, 2004; Giannakopoulos et al., 2005; Xoplaki et al., 2005; et al. (2004), climate change is today one of the most serious Luterbacher et al., 2007). By the 2020s, under high CO2 scen- threats to biodiversity and ‘on the basis of mid-range climate arios, summer in southern Europe will be as hot as or hotter warming scenarios for 2050, 15–37% of species in our than the 2003 summer (Luterbacher et al., 2004). sample of regions and taxa will be “committed to extinction”’. Font et al. (2007) have documented that the exceptional As a miniature ocean (0.82% of the world’s ocean surface) winter of 2005 affected the deep-water thermohaline proper- with fast turnover time (40–50 years) the semi-enclosed ties of the north-western Mediterranean. Similarly, in the Mediterranean, which accounts for 8% to 9% of global Aegean Sea, it is likely that changes in the freshwater budget marine biodiversity and contains numerous endemic species could have modified seawater salinity and altered water-mass (Bianchi & Morri, 2000), is likely to rapidly respond to exter- circulation (Roether et al., 1996; Theocharis et al., 1999). nal forcing like climate change (Be´thoux & Gentili, 1996; Climate variation—including extreme events—is not necess- Monaco, 1998; Be´thoux et al., 1999; Turley, 1999; arily a result of directional climate change. However, climate Simmonds & Nunny, 2002; Giorgi, 2006). change is likely to create more extreme meteorological The aim of this review is to illustrate the links existing events (e.g. storms and drought), and these can alter the bio- between biodiversity and climate and demonstrate how anthro- chemical and physical properties of seawater (Poumade`re pogenic greenhouse warming can affect the Mediterranean et al., 2005; Olita et al., 2007). marine ecosystems including large predators such as cetaceans. Itillustratestheimpactsofglobal warming on abiotic and sea level rise biotic systems in the Mediterranean Sea and stresses the neces- Ecologically and socio-economically rich coastal marine sity to consider climate change as a major issue. systems are under threat from anthropogenic global warming (IPCC, 2001a; Roessig et al., 2004). Sea level rise resulting from climate change-related processes such as sea- DISCUSSION water thermal expansion or glacier melt will trigger coastal flooding (Paskoff, 2001). According to Tsimplis & Rixen Influence of climate change on Mediterranean (2002) from 1993 onwards, the warming of surface waters in the Eastern Mediterranean has caused sea level rise. In abiotic parameters the last century, sea level has risen by 0.10 to 0.20 m around By influencing climatic features (e.g. atmospheric tempera- Europe (IPCC, 2001a) and an increase of sea level of 3.3 cm ture, precipitation budget and extreme meteorological in 11 years (0.3 cm per year) was recorded at the oceano- events), global warming is likely to affect the chemical and graphic and meteorological station at l’Estartit in north-east physical properties of Mediterranean waters. Spain (Salat & Pascual, 2002). Between 1990 and 2100, the predicted trend for sea level rise is 2.2 to 4.4 times higher than the rate recorded in the 20th Century (IPCC, 2001a). Present and future trends of Mediterranean Consequently, some Mediterranean coastal areas and wet- climate and hydrology lands, such as the Camargue, are threatened (Nicholls & Hoozemans, 1996; Pfeifle et al., 2004). At the beginning of the 20th Century, the temperature and salinity of the Western Mediterranean Deep Water (WMDW) was practically constant (Lacombe et al., 1985; Be´thoux et al., atmospheric change and climate impacts in 1990). Over the last few decades, numerous studies have indi- the mediterranean cated a correlation between climate patterns and changes in sea- Anthropogenic climate change tends to influence large atmos- water properties, with general increases in both temperature pheric patterns like the North Atlantic Oscillation (NAO) and and salinity over the entire Mediterranean (Lacombe et al., the El Nin˜o Southern Oscillation (ENSO) (Timmermann 1985; Be´thoux et al., 1990; Leaman & Schott, 1991; Rohling & et al., 1999; Visbeck et al.,2001;Wu¨rsig et al., 2002; Cohen & Bryden, 1992; Francour et al., 1994; Sparnocchia et al.,1994; Barlow, 2005). Global warming is likely to contribute to the posi- Astraldi et al., 1995; Graham, 1995; Be´thoux & Gentili, 1996; tive phase of the NAO, which was responsible for warmer Be´thoux et al., 1998, Krahmann & Schott, 1998; Duarte et al., winters in Europe in the last 10–20 years (Hurrell, 1996; 1999; Danovaro et al., 2001; Astraldi et al., 2002; Fuda & Visbeck et al., 2001; Gillet et al., 2003; Cohen & Barlow, 2005) Millot, 2002; Gertman & Hecht, 2002; Goffart et al.,2002; and El Nin˜o conditions may be more frequent with global Lascaratos et al., 2002; Lopez-Jurado, 2002; Manca et al., warming (Timmermann et al., 1999; Wu¨rsig et al., 2002). 2002; Prieur, 2002; Salat & Pascual, 2002; Vargas-Yanes et al., The Mediterranean climate is influenced by large-scale 2002; Vilibic, 2002; Walther et al., 2002; Rixen et al.,2005;Li atmospheric circulation systems including the inter-decadal et al., 2006; Millot et al., 2006; Somot et al., 2006). It is now NAO and ENSO (Hurrell, 1995; Raicich et al., 2001; assumed that the WMDW has become warmer and saltier in Mariotti et al., 2002; Lionello & Sanna, 2005). Changes in response to anthropogenic greenhouse effects and changes in Mediterranean seawater temperature and salinity have been the freshwater budget (Be´thoux et al., 1990; Rohling & associated with the NAO (e.g. Be´thoux & Gentili, 1999; Bryden, 1992; Graham, 1995; Be´thoux et al., 1998). Be´thoux et al., 1999; Tsimplis et al., 2006). In fact, the environmental threats to mediterranean cetaceans 181

NAO influences Mediterranean precipitation and river potential effects of climate discharges (nutrient inputs), which tend to be lower during change-induced environmental variations positive NAO episodes (Lloret et al., 2001; Struglia et al., on mediterranean marine species 2004; Xoplaki et al., 2004, 2006). According to Tsimplis & 1. Climate impacts on marine fauna Josey (2001), NAO is also influencing sea level change in The global warming-induced alteration of precipitation, temp- the Mediterranean Sea. Mariotti et al. (2002) have also ident- erature, CO concentration and wind patterns will result in a ified a link between ENSO and Mediterranean autumn 2 cascade of changes in the physical (e.g. vertical stability of rainfall anomalies. The relationship between NAO, ENSO the water column, upwelling regimes, water mass formation and climate change is an important issue and requires and circulation, current patterns), chemical (e.g. seawater further study. pH, salinity, nutrient ratios) and biological (e.g. species phenol- ogy, recruitment, physiology, distribution, abundance, diver- sity, productivity) properties of marine systems (Figure 1) Climate change impacts on Mediterranean (e.g. Bianchi, 1997; Walther et al., 2002). Temperature marine biodiversity and ecosystems anomalies, even over a short period of time, can significantly affect Mediterranean ecosystems and biological diversity Evidence has demonstrated that climate variation and changes (Walther et al., 2002; Anado´n et al., 2007). When a keystone in the properties of ecological systems are strongly correlated species is affected, even by a slight change in climate, the com- (e.g. Orr et al., 1992; Astraldi et al., 1995; Bombace, 2001; position and diversity of marine communities can be disrupted Hannah et al., 2002; Walther et al., 2002; Duffy, 2003; Root (Sanford, 1999). The structure, distribution and phenology of et al., 2003; Roessig et al., 2004) and that global warming is Mediterranean plankton communities, which are at the base already affecting numerous marine species throughout the of the food chain and which strongly depend on hydroclimatic world (Table 1). In the Mediterranean, it is now acknowledged factors and nutrient ratios, are experiencing change (Velsch, that climate change-induced temperature variations have 1997; Turley, 1999; Licandro & Ibanez, 2000; Be´thoux et al., altered biological patterns and biodiversity (e.g. Francour 2002; Fernandez de Puelles et al., 2004; Mercado et al., 2005; et al., 1994; Turley, 1999; Bianchi & Morri, 2000). Molinero et al., 2005a, b; Molinero et al., 2007; Voarino, 2006). For instance, high positive anomalies in water tempera- ture during the 1980s resulted in jellyfish and a drop in abun- Table 1. Examples of species affected by climate change induced-factors dance of copepods (Molinero et al., 2005a). around the world (Mediterranean Sea excluded). Climate has long been recognized as one of the most critical Areas Examples of affected species factors influencing Mediterranean resource variability (Garcia & Palomera, 1996). For instance, the most essential environ- North-east Atlantic † Marine flora: Ducrotoy (1999) mental mechanisms controlling the growth, abundance, † Plankton: Southward et al. (1995); Fromentin distribution, composition, diversity and recruitment success & Planque (1996); Nehring (1998); Planque of Mediterranean species, such as anchovy (Engraulis encrasi- & Taylor (1998); Heath et al. (1999); Edwards et al. (2001); Beare et al. (2002); colus) or sardine (Sardinella aurita and Sardina pilchardus), Beaugrand et al. (2002); Edwards et al. include: regional temperature variations, riverine inputs and (2002); Sims & Reid (2002); Beaugrand wind-induced mixing, which influence sea surface temperature (2003); Beaugrand & Reid (2003); Edwards & and salinity; hydrographical features (e.g. oceanic fronts, water Beaugrand (2007); Greve (2007); Pitois & column stability, upwelling zones, sea state); and nutrient Fox (2007); Slagstad et al. (2007); Voss et al. enrichment and planktonic production (e.g. Cury & Roy, (2007) 1989; Sabates & Gili, 1991; Garcia & Palomera, 1996; Regner, † Intertidal and benthic organisms: Southward 1996; Agostini & Bakun, 2002; Lafuente et al., 2002; LLoret et al. (1995); Kroncke et al. (1998) et al., 2004; Lafuente et al., 2005; Basilone et al., 2006; † Fish: Scarnecchia (1984); Svendsen (1995); Sabates et al., 2006). Alheit & Hagen (1997); Southward et al. (1988); Sims & Reid (2002); Beaugrand & The Mediterranean Sea is considered oligotrophic because Reid (2003); Beaugrand et al. (2003); Clark of its low nutrient input from rivers and the nutrient depleted et al. (2003); Genner et al. (2004); Sims et al. Atlantic water inflow through the Strait of Gibraltar (e.g. (2004); Brander (2005); Cotton et al. (2005); Estrada, 1996; Turley, 1999; Zenetos et al., 2002). Winter ver- Perry et al. (2005); Rose (2005); Sissener & tical mixing, coastal upwelling and river runoff are the mech- Bjorndal (2005); Po¨rtner & Knust (2007); anisms of nutrient input into the euphotic zone, on which Stenevik & Sundby (2007); Voss et al. (2007) marine productivity is dependent. Ocean physical processes † Cephalopods: Sims et al. (2001) such as upwelling phenomena have a major influence on the Tropical Atlantic † Plankton: Piontkovski & Castellani (2007) distribution of primary production, through ascending move- ments of nutrient-rich deep water into the euphotic zone. Antarctic Peninsula † Plankton: Moline et al. (2004) Therefore, by altering oceanic features, climate change may Bay of Biscay † Fish: Poulard & Blanchard (2005) affect nutrient availability. Increasing temperatures are likely to trigger stronger † North Pacific Ocean Plankton: Roemmich & McGowan (1995); thermal stratification and deepen the thermocline, which Brodeur et al. (1999); Batten (2007) could prevent or modify the mixing of water-masses, and Worldwide † Coral reefs: Sebens (1994); Sheppard (1999); cold and nutrient-rich deep waters upwelling (Roemmich & Spalding et al. (2003) McGowan, 1995). Some areas like the southern Adriatic Sea, where local winter climatic conditions (e.g. winter heat 182 d.d. gambaiani et al.

Fig. 1. Main greenhouse gases-related factors influencing marine organisms.

losses and precipitation) strongly influence nutrient avail- biological pump of carbon into the deep ocean and strongly ability and plankton production (Gacic, 2002), may be influence fish recruitment could alter the ecosystem function- especially vulnerable to climate change. ing (Ohman & Hirche, 2001; Molinero et al., 2005b). Extreme meteorological events (e.g. storms and flooding Nutrient inputs from the Rhoˆne and Ebre rivers, episodes) and sea level rise will release abundant terrestrial combined with strong wind mixing, make the north-western suspended solids and pollutants into the marine environ- Mediterranean highly productive (Lloret et al., 2001). As ment, which may negatively affect coastal biocenoses. previously noted, climate change is expected to induce Fragile biotopes of the endemic Posidonia oceanica have positive NAO episodes with less precipitation and runoff. been shown to be vulnerable to physical and chemical Consequently, the recruitment of north-western Mediterranean damage from meteorological events such as these (Orr fish, which strongly depend on riverine nutrient supply, may be et al., 1992; Bombace, 2001). As these sea grass meadows negatively affected by climate change (Lloret et al., 2001). represent a spawning and nursery habitat for numerous In certain areas some organisms survive under specific species and play a major ecological role, their disappearance temperature conditions and cannot adapt themselves or would to expected to have significant consequences for move when the environmental conditions change (e.g. coastal ecosystems (Francour, 1997). Kenney, 1990; MacGarvin & Simmonds, 1996; Hughes, By influencing the north-western Mediterranean climate 2000). Dispersal limitation can limit the response of marine variability (Pozo-Vasquez et al., 2001; Gasparini & Astraldi, animals such as benthic organisms that could sometimes be 2002; Rixen et al., 2005), the NAO affects the local species com- unable to successfully migrate toward more suitable environ- position of planktonic copepods, such as the two dominant ments (problems would include long distances or travelling copepods of the north-west Meditarranean, Centropages against strong currents) (Hiscock et al., 2004). typicus and Temora stylifera (Molinero et al., 2005a, b). By affecting the physiology of marine organisms, global The high positive NAO episode that caused positive temp- warming could impact the performance and survival of erature anomalies in the 1980s in the western Mediterranean those organisms that live close to their thermal tolerance or Sea induced a jellyfish bloom and resulted in a significant situated at the northern or southern limit of their distribution diminishment of copepod abundance (Molinero et al., (Laubier, 2001; Hochachka & Somero, 2002; Somero, 2002; 2005a). This event, which is likely to have been encouraged Poulard & Blanchard, 2005). Similarly, some larval and by global warming, led to high abundance of Centropages young benthic stages of some organisms are more sensitive typicus at the expense of Temora stylifera (Molinero et al., to temperature than adults (Foster, 1971; Pechenik, 1989). 2005b). Jellyfish, which feed on fish larvae, eggs and copepods, According to Bella Galil in Cheviron (2007), deep-water can strongly affect plankton communities (Mills, 1995). organisms that live in constant temperatures (138C) and Changes in planktonic copepods, which affect the fluxes of that are not used to seasonal temperature variation will be vul- matter and energy in the marine ecosystem, supply a nerable to climate change. environmental threats to mediterranean cetaceans 183

2. Climate impacts on marine flora In the Mediterranean, over the past three decades, increas- Climate change is likely to affect phytoplankton composition by ing water temperature has been observed in the Ligurian Sea affecting nutrient concentrations and ratios. Over the last two (Be´thoux et al., 1990; Astraldi et al., 1995), which is one of decades, in the north-western Mediterranean, meteorological the coldest zones of the Mediterranean. This phenomenon anomalies (e.g. warmer water, decreased salinity, longer periods encouraged warm-water species to shift their ranges northward of sunshine and lower wind stress) have affected water column and settle in the Ligurian waters where they were formerly rare stability and reduced nutrient replenishment into the euphotic or absent (Bianchi & Morri, 1993, 1994; Francour et al., 1994; zone (Goffart et al., 2002). This event decreased silicon avail- Astraldi et al., 1995; Morri & Bianchi, 2001). ability, which in turn triggered a reduction of diatom abundance Consequently, warm-water species like the ornate wrasse and a shift toward non-siliceous species, such as flagellates and (Thalassoma pavo) colonized and established large and stable dinoflagellates (Turley, 1999; Be´thoux et al., 2002; Goffart et al., populations in the north-western Mediterranean (Bianchi & 2002). By modifying the composition of phytoplankton commu- Morri, 1994; Bombace, 2001; Vacchi et al., 2001). Other thermo- nities, climate change could then seriously alter nutrient cycling philic species like the grey triggerfish (Balistes carolinensis), and food web dynamics (Litchman et al., 2006). Mediterranean parrotfish (Sparisoma cretense), round sardine Warmer temperatures trigger changes in the timing of (Sardinella aurita), bluefish (Pomatomus saltatrix), Senegalese plankton blooms, resulting in a temporal mismatch between sole (Solea senegalensis), dusky grouper (Epinephelus margina- primary production and higher trophic levels of the food tus), bastard grunt (Pomadasys incisus), European barracuda web (Edwards & Richardson, 2004; Hiscock et al., 2004). In (Sphyraena sphyraena), slackskin blaasop (Sphoeroides cutaneus), north-western European estuaries, for example, a long-term the coral Astroides calycularis and groupers of the genus data set (1973–2001), showed an increase of spring tempera- Epinephelus have been frequently recorded amongst the ‘cold tures of 0.078Cyr21, which resulted in the earlier spawning of biota’ of the northern Mediterranean (Vacchi & Cau, 1986; a bivalve, Macoma balthica, but did not affect phytoplankton Serena & Silvestri, 1996; Relini & Orsi Relini, 1997; Dulcic et al., blooms, causing a divergence in timing between larval bivalve 1999; Louisy & Culioli, 1999; Dulcic & Grbec, 2000; Guidetti production and prey (Philippart et al., 2003). In addition, & Boero, 2001; Dulcic et al., 2005, 2006; Athanassios & rising seawater temperatures advance the onset of crustacean Antonopoulou, 2006). reproduction and enhance shrimp predation pressure on The recent northward expansion of tropical groupers, like vulnerable juvenile spat that leads to low recruitment the white grouper (Epinephelus aeneus) and dusky grouper success of the spat (Philippart et al., 2003). (Epinephelus marginatus), may result from the warming of Mediterranean waters (Francour et al., 1994; Dulcic & Lipej, migration of species and increased number 1997; Zabala et al., 1997). Since groupers are top carnivores and among the bigger coastal fish species, their successful of exotic thermophilic species in the colonization is likely to affect the ecology of endemic species mediterranean sea and influence local fisheries (Glamuzina, 1999). The presence of exotic species in the Mediterranean has Similarly, in the Adriatic Sea, the presence of thermophilic resulted from the combination of environmental factors and species of fish and zooplankton, that were formerly uncom- human activities like the opening of the Suez Canal in 1869. mon or absent in this zone, has increased in the past 30 Today, more than 500 alien species are recorded in years (Dulcic & Grbec, 2000; Kamburska & Fonda-Umani, the Mediterranean Sea and their geographical ranges are 2006). These observations have been correlated with seawater increasing, as is the rate of increase of new alien species warming and salinity changes occurring since 1988 (Francour being identified (Galil & Zenetos, 2002; Zenetos et al., 2003; et al., 1994; Russo et al., 2002) and, thus, global warming is Harmelin-Vivien et al., 2005; CIESM, 2005; Galil, 2007). thought to be responsible for the observed faunal changes in Climate change, combined with the establishment of exotic this region (Dulcic et al., 1999; Dulcic & Grbec, 2000; species has led to the ‘tropicalization’ of the Mediterranean Parenti & Bressi, 2001). (Bianchi, 2007). The increase of alien species can cause The northward migration of warm water species will induce endemic species to rapidly decline in abundance and be dis- species competition for existing niches (IPCC, 2001a) and placed (Galil & Zenetos, 2002; Zenetos et al., 2002; Galil, thermophilic species are likely to increase in abundance at 2007). Such phenomena can alter the infra-littoral commu- the expense of cold-water species (Beaugrand et al., 2002; nities and induce ecological impacts such as local population Galil & Zenetos, 2002; Hiscock et al., 2004). Marine bioinva- decline and extirpation, reduction of genetic diversity in sions, which are considered to be ‘biological pollution’ native species, foodweb alterations, loss of habitat functions, (Bouderesque & Verlaque, 2002; Elliot, 2003) have altered processes and structure, increase in the risk of extinction marine ecosystems (e.g. competition with indigenous species, and biotic homogenization (Ricciardi, 2004; Galil, 2007). food web shifts) and are considered as one of the most In the North Atlantic Ocean, the observed northward intense and damaging anthropogenic impacts (Harris & migration of species (250 km per decade), resulting from a Tyrrell, 2001; Frank et al., 2005; Galil et al., 2007). The minor increase in temperature, is likely to be linked to Mediterranean Sea is one of the most impacted seas of the global warming (Parmesan & Yohe, 2003; EEA, 2004; world in terms of biological invasion (Galil, 2007) and Oviatt, 2004). Perry et al. (2005) have also demonstrated climate change is likely to facilitate invasion of thermophilic that, in the North Sea, recent increases in sea temperature alien species causing irreversible impacts on native populations have led to nearly two-thirds of fish species (exploited and (Carlton, 2000; Stachowicz et al., 2002b; Gritti et al., 2006; Galil non-exploited) shifting in mean latitude or depth or both et al., 2007; Occhipinti-Ambrogi, 2007). According to Bella over 25 years. For species with northerly or southerly range Galil (in Cheviron, 2007) alien species like the bivalve margins in the North Sea, half showed boundary shifts with Brachidontes pharaonis or the jellyfish Rhopilema nomadica, warming, and all but one species shifted northward. both belonging to the 100 ‘worst invasives’ species, could 184 d.d. gambaiani et al.

lead to the extinction of numerous native Mediterranean bryozoans, scleractinian corals and zoanthids (Cerrano et al., species. Photophilic subtidal macrophyte assemblages appear 2000; Perez et al., 2000; Romano et al., 2000; Garrabou et al., particularly vulnerable to invasions of exotic algal species 2001). According to Occhipinti-Ambrogi (2007), niches result- such as Caulerpa taxifolia (Streftaris & Zenetos, 2006). ing from mass mortality events become available for new inva- In addition to northward migration, bathymetric displace- sive colonizers. ments occur among populations of invasive and endemic Variations of the nutrient load and seawater properties (e.g. species (Galil & Zenetos, 2002). This is the case for the indigen- temperature increase) can lead to coastal eutrophication and ous red mullet (Mullus barbatus)andhake(Merluccius merluc- algal bloom (Degobbis et al., 2000). This phenomenon particu- cius) with both moving into deeper and cooler waters due to larly concerns the shallow northern Adriatic waters (UNEP, their respective warm-water competitors: the goldband goatfish 1996; Degobbis et al., 2000). By increasing seawater stratifica- (Upeneus moluccensis) and brushtooth lizardfish (Saurida tion, meteorological anomalies regularly cause bottom water undosquamis) (Oren, 1957). Similarly, in the north-western anoxia and red tide events in this region, provoking mass mor- Mediterranean, the endemic spottail mantis shrimp (Squilla tality episodes of fish and benthic organisms (Degobbis et al., mantis) is usually observed in deeper waters (70–80 m) rather 2000; Anado´n et al., 2007). Eutrophication events occurred fre- than the thermophilic Red Sea mantis shrimp (Oratosquilla quently during the second half of the 20th Century (Degobbis massavensis) (10–25 m) (Galil & Zenetos, 2002). et al., 2000) and the intensity, frequency and geographical The warming of the Mediterranean waters may modify expansion of algal blooms became a growing concern since species’ migration periods causing changes in the trophic the 1970s in this area (Justic, 1987). According to Boero webs. For example, Bombace (2001), has documented that in (1996), increases in jellyfish (Pelagia noctiluca and Aurelia the last few decades, the amberjack (Seriola dumerilii)and aurita), salps (Thaliacea), harmful algal blooms and red tides bluefin tuna (Thunnnus thynnus) appear to stay longer (until were all promoted by abnormal meteorological and oceano- mid-winter instead of autumn) in the northern and central graphic changes occurring since 1988 in the Adriatic Sea. Mediterranean before migrating toward their winter territories. During the last decades, the collapse of sprat (Sprattus Although there are presently no marine species extinctions sprattus) and anchovy (Engraulis encrasicolus) stocks in the that have been correlated with global warming, today, species Adriatic affected the species stocks of the entire Mediterranean such as the Mediterranean mysid Hemimysis speluncola and was associated with the decrease of surface temperature are regarded as threatened (Chevaldonne´ & Lejeusne, 2003; resulting from climatic anomalies (Regner, 1996; Salat, 1996; Lejeusne, 2005; Lejeusne & Chevaldonne´, 2005; Harley Bombace, 2001; Azzali et al., 2002). Similarly, during the 2001 et al., 2006). Furthermore, Chevaldonne´ & Lejeusne (2003) winter, climate-induced low surface temperatures led to the showed that the endemic cave-dwelling invertebrate decline of sardines (Sardinella aurita) and phytoplankton Hemimysis speluncola which was previously abundant in the blooms (Guidetti et al.,2002). north-western Mediterranean, has been replaced by a warm- By altering oceanographic properties like seawater temp- water species (Hemimysis margalefi). erature, salinity, water transparency and deep water oxygen Areas of high endemic biodiversity are likely to be less saturation, climate change could impact the entire Adriatic subject to non-indigenous species invasion (Kennedy et al., Sea ecosystem (Zore-Armanda et al., 1987; Zore-Armanda, 2002; Stachowicz et al., 2002a; Duffy, 2003). Therefore, 1991; Russo et al., 2002). sparse and declining populations such as Posidonia meadows Global warming may promote the development of toxic are more vulnerable to be overgrown and replaced by invasive dinoflagellates like Gymnodinium catenatum which is respon- macroalgae like Caulerpa taxifolia and Caulerpa racemosa sible for frequent Mediterranean toxic events, causes paralytic (Meinesz et al., 2001; Bouderesque & Verlaque, 2002; shellfish poisoning (PSP) and may alter the marine ecosystem Peirano et al., 2005). Climate change could be one of the (Garce´s et al., 2000; Taleb et al., 2001; Vila et al., 2001; Calbet factors responsible for the decline of Posidonia and for the et al., 2002; Gomez, 2003). According to Calbet et al. (2002), expansion of Caulerpa (Komatsu et al., 1997; Raniello et al., the occurrence of Gymnodinium catenatum (toxic dinoflagel- 2004; Peirano et al., 2005; Ruitton et al., 2005). late) in the Alboran Sea reduced the grazing impact of meso- Furthermore, some cold-water species like the small zooplankton on the microbial communities and may have euphausid species Meganyctiphanes norvegica, which is situ- altered the Mediterranean pelagic food web. ated at the northern limit of its ecological tolerance, would Gomez & Claustre (2003) and Polat (2004) suggest that the be more vulnerable to invasion. This is especially true for presence of new warm-water dinoflagellate species in the many taxa in the eastern Mediterranean, leaving this region Mediterranean Sea, like Asterodinium libanum, Asterodinium more exposed to invasion (Galil & Zenetos, 2002). gracile and Citharistes regius is likely to be associated with the warming of Mediterranean waters. A species of a similar large mass-mortality events and decline in genus, Ostreopsis armata, was recently observed (summer 2005 and 2006) in the Ligurian Sea where it repeatedly species abundance and diversity associated bloomed and apparently triggered respiratory diseases in with temperature anomalies humans (details in Occhipinti-Ambrogi, 2007). Temperature anomalies have led to several mass-mortality Meteorological anomalies can significantly alter ecosystems episodes in the Mediterranean Sea. In 1999, successive heat and cause mass mortality episodes. In the eastern Mediterranean waves with consequent peaks in water temperature and a dee- Sea, a climatic event, called the Eastern Mediterranean pening of the thermocline caused a mass-mortality event of 28 Transient (EMT), was correlated with local meteorological invertebrate species in the north-western Mediterranean anomalies (reduced precipitation, change in wind patterns (Cerrano et al., 2000; Perez et al., 2000; Romano et al., 2000; and cold winters) (Klein et al., 1999) and resulted in a Laubier et al., 2003). This event affected benthic organisms drastic alteration of faunal abundance and diversity such as gorgonians, sponges, cnidarians, bivalves, ascidians, (Danovaro et al., 2001). By modifying the physico-chemical environmental threats to mediterranean cetaceans 185 characteristics of the deep waters, this temperature shift sig- celled protists to reef-building corals—and across all CaCO nificantly and rapidly affected deep-sea nematode diversity mineral phases. Cephalopods may also be particularly sensitive (Danovaro et al., 2004). Following the 1994–1995 period, and this is described in Table 2 at the end of this paper. when the temperature recovered, only some marine faunal According to a study recently carried out by The Royal species recovered (Danovaro et al., 2004). These observations Society (2005), the plankton calcification process could give us a better vision of the potential large-scale consequences become very limited and a large portion of marine life could of global warming. then disappear by 2100. The seawater acidification phenomenon, which is respon- increase of diseases and pathogens sible for nanism (a genetic anomaly resulting in short Since warmer temperatures are known to favour the presence stature) and malformation symptoms in several phytoplankton of pathogens, epidemiological outbreaks are likely to become species (Feely et al., 2004), can also have an impact on the more severe and frequent with the warming of Mediterranean reproductive patterns of fish (Stanley, 1984). waters (Gantzer et al., 1998; Harvell et al., 1999, 2002; According to Caldeira & Wickett (2003) and Feely et al. Marcogliese, 2001; Drake et al., 2007). According to CIESM (2004), the expected change in pH will be greater than any (2004) global warming could lead to the development of tro- other pH variations observed in the fossil record over the last pical and subtropical pathogens across the Mediterranean Sea. 200–300 million years. Manipulative experiments showed Along the Ligurian Sea, the massive development of a cyano- that a three-month reduction of pH by 0.7-unit reduced bacterium combined with warm water, caused several popu- mussel metabolism and growth (Michaelidis et al., 2005). lations of the zoanthid Parazoanthus axinellae to decline Similarly, a six-month, 0.03-unit pH reduction, which corre- since 2000 (Cerrano et al., 2006). This species has been sponds to a 200-ppm increase in atmospheric CO2, lowered replaced by an incrusting thermophilic sponge (Crambe gastropod and sea urchin growth and survival (Shirayama & crambe) which quickly colonized the niche deserted by Thornton, 2005). A study by Alvarez et al. (2005) shows that, Parazoanthus (Cerrano et al., 2006). although ocean uptake of carbon dioxide for the Similarly, the thermophilic bacteria Vibrio shiloi, was Mediterranean appears small in terms of global ocean uptake, involved in the mortality episode of the Mediterranean coral ‘the impact on local carbonate chemistry will be large.’ Oculina patagonica (Kushmaro et al., 1998) and the 1999 In addition to this process of ocean acidification, changes mass mortality event in the Ligurian Sea was induced by the in marine ecosystems due to global warming (e.g. water temp- combination of a temperature shift with the growth of oppor- erature, algal bloom enhancement and water colour) might tunistic warm-water pathogens (Cerrano et al., 2000). affect the visual sensitivity of fish (spectral sensitivity) Simmonds & Mayer (1997) provided a tentative link between (Archer et al., 2001). reduced nutrient input to the western Mediterranean basin Several points discussed in this section are likely to be relevant (resulting from reduced rain fall) and the initiation of the to a broad variety of marine species including key species in striped dolphin (Stenella coeruleoalba) mass mortality in 1990. Mediterranean ecosystems, with significant consequences for This is discussed further below. foodweb structures, marine system functions and ecosystem equilibrium (Petchey et al., 1999; Sanford, 1999; Schiel et al., other expected changes 2004). Such foodweb alterations are likely to produce significant Increased concentration of atmospheric anthropogenic CO2 cascade effects on marine biodiversity including impacts on and climate change is likely to alter marine ecosystems in species of higher trophic levels, such as cetaceans. several other ways. As temperature increases, the oxygen solu- bility decreases and fish metabolism accelerates (Green & Carritt, 1967; Po¨rtner & Knust, 2007). Since the demand for Climate change impacts on cetaceans oxygen and food will be enhanced in order to support higher The distribution of cetaceans, which have a major influence on metabolic rates, the decreased concentration of dissolved marine community function and structure (e.g. Katona & oxygen will thus affect fish growth and breeding capacity, and Whitehead, 1988; Bowen, 1997; Jones et al., 1998), is closely could lead to the extinction or migration (to cooler waters) of related to environmental parameters such as oceanographic some fish species (Po¨rtner & Knust, 2007). Furthermore, features and food availability (Millot & Taupier-Letage, enhanced fish metabolic rates and food consumption resulting 2004). According to MacGarvin & Simmonds (1996), they from higher seawater temperatures may increase fish pollutant are not likely to be able to adapt to rapid shifts in temperatures uptake (e.g. mercury) which will then be transferred into higher and environmental conditions, and climate change may rep- levels of the food chain (Harris & Bodaly, 1998). resent the most serious long-term threat to cetaceans In addition, the rise of atmospheric CO could increase the 2 (Burns, 2001). Although lower trophic levels are the most acidity of seawater and therefore reduce the saturation state of likely to be altered, cetaceans could be affected by global CaCO species in the oceans, namely calcite and aragonite 3 warming in a variety of ways (Figure 2; Table 2) (e.g. (Caldeira & Wickett, 2003; Orr et al., 2005). The marine Reeves, 1991; Fisher et al., 1994; Agardy, 1996; IWC, 1997; species most likely to be affected by this acidification will be European Community, 1999; Hardwood, 2001; Simmonds & small and thin-shelled organisms that use CaCO such as cal- 3 Nunny, 2002; Gambaiani et al., 2005; Learmonth et al., cifying plankton (e.g. coccolithophores), coralline algae, ptero- 2006; Simmonds & Isaac, 2007). pod molluscs and coral polyps (e.g. reef-building scleractinian corals) (Kleypas et al., 1999; Riebesell et al., 2000; Feely et al., 2004). By reducing the level of calcium carbonate saturation, change in food supply ocean acidification will affect the process of calcification of The distribution, abundance and migration of cetaceans is some marine key organisms. Feely et al. (2004) argue that the strongly influenced by prey availability (e.g. Kenney et al., calcification rate of multiple taxa will be affected, from single 1996) and cetaceans which are confined in restricted habitat, 186 d.d. gambaiani et al.

Fig. 2. Overview of the potential impacts of global warming on marine life, including cetaceans (from Gambaiani et al., 2005).

withlimitedranges,arelikelytobemostvulnerabletoclimate and energy foraging, which could have drastic consequences change (e.g. Learmonth et al., 2006; Simmonds & Isaac, 2007). on their health and could affect their immune systems Change in key prey species distribution is the main driving (Northridge, 1984; Shane, 1990; Bra¨ger, 1993; Smith & factor defining geographical range and habitat preference in Whitehead, 1993; Agardy, 1996; Stern, 1996; Bearzi, 2002). A cetaceans (e.g. Evans, 1971; Wells et al., 1990; Hanson & high proportion of time and effort devoted to feeding-related Defran, 1993; Simmonds, 1994; Agardy, 1996; Maze & activities was recorded in Mediterranean bottlenose dolphins Wu¨rsig, 1999). in the northern Adriatic Sea as a response to environmental For instance, in the eastern North Pacific Ocean, an changes and reduced prey availability (Politi, 1998; Bearzi increase in seawater temperature combined with a change in et al., 1999). Consequently, the time dedicated to socializing oceanographic conditions is thought to have led to the death and breeding is reduced, with negative consequences on ceta- of hundreds of grey whales (Eschrichtius robustus) as the cean reproductive success (Valiela, 1995; Bearzi, 2002). The result of a decline in their prey species (Grebmeier & fact that climate change impacts can lead to reduced prey avail- Dunton, 2000; Moore et al., 2003; Gulland et al., 2005). In ability and subsequently affect the health, physical strength and the Mediterranean Sea, the decline of several cetacean popu- abundance of cetacean populations, has been observed in bottle- lations has been associated with the reduction of prey nose dolphins in the eastern Ionian Sea (Politi et al., 2000; Politi resources (Perrin, 1989; UNEP/IUCN, 1994; Reeves et al., & Bearzi, 2004). 2003; Reeves & Notarbartolo di Sciara, 2006). Aguilar & Raga (1993) and Simmonds & Mayer (1997) have Odontocete prey species such as pilchard (Sardina pilchar- suggested that the mass mortality of thousands of striped dol- dus) are affected by climate change (Southward et al., 1988; phins (Stenella coeruleoalba) during the 1990–1992 morbilli- Garcia & Palomera, 1996; Regner, 1996; Bearzi et al., 2003). virus epizootic might have been caused by the unusual warm Pilchards have been shown to be a key prey species for and dry winter of 1989–1990 that led to abnormally warm common dolphins off the Portuguese coast (Silva, 1999). In water temperatures and low rainfall. This phenomenon addition, cephalopods, which represent the main food resulted in reduced nutrient input into the eastern supply for numerous Mediterranean cetacean species (Wurtz Mediterranean and thus low productivity (Simmonds & & Marrale, 1991; Bompar, 2000; Bearzi et al., 2003) seem to Mayer, 1997). This led to the decline of the dolphin’s be particularly vulnerable to environmental changes including common prey and explains why many of the dolphin carcasses pH and temperature (Sims et al., 2001; Pierce & Boyle 2003; showed depleted body fat reserves (Aguilar et al., 1991). Arkhipkin et al., 2004) (Table 2). Similarly, during a survey carried out in Corsica, Dhermain In the Adriatic Sea, climatic shifts are suggested to have (2003) observed fewer bottlenose dolphins in coastal waters altered the distribution of the key prey species of common than usual. The excessively hot 2003 summer, which resulted dolphins (Delphinus delphis) and bottlenose dolphins in abnormally high coastal water temperatures, could explain (Tursiops truncatus) (Blanco et al., 2001; Bearzi et al., 2003). the migration of bottlenose dolphins toward the open sea. In particular, the climate-induced increase in abundance of Such exceptional meteorological events may illustrate how thermophilic species such as round sardinella and jellyfish an increase of temperature could impact on cetaceans. may have caused the European anchovy (Engraulis encrasico- Moreover, global warming is likely to encourage the lus) population to decrease (Regner, 1996). spreading of viruses and pathogens and may promote Shifts in prey species availability may force cetaceans to epizootic events like morbillivirus infections (Agardy, 1996), change their feeding strategies and spend more time which have also been identified in the endangered environmental threats to mediterranean cetaceans 187

Table 2. Impact of climate changes on cetaceans around the world.

Global Impacts on marine mammals with some examples of species most likely to be, or, already affected warming- induced changes

Seawater Directly affected: † North-east Atlantic fin whales (Lockyer, 1986) temperature † Endangered, young animals, species with low mobility, † Humpback whales (Wiley & Clapham, 1993) restricted distribution, low thermal tolerance like vaquitas † Harbour porpoises (Phocoena phocoena) in the Bay of Fundy (Phocoena sinus) in the Gulf of Mexico; Arctic bowhead (Read & Gaskin, 1990) whales (Balaena mysticetus); Arctic belugas (Delphinapterus Effects on reduced prey availability leading to: leucas); narwhals (Monodon monoceros); finless porpoises † Increased vulnerability to disease particularly for species at (Neophocaena phocaenoides), tropical dolphins, humpback whales (Megaptera novaeangliae) in the Indian Ocean, the limit of their thermal tolerance (Wu¨rsig et al., 2002; Lafferty et al., 2004) Mediterraneanfinwhales(Balaenoptera physalus)and † river dolphins (IWC, 1997; Bannister, 2002; Wu¨rsig et al., Increase of mass mortality events like morbillivirus that affected Mediterranean striped dolphins (Stenella 2002; Simmonds, 2004; COSEWIC, 2005; Laidre & coeruleoalba) (Aguilar & Raga, 1993; Cebrian, 1995; Harvell Heide-Jorgensen, 2005; Robinson et al., 2005; Learmonth et al., 2006; Simmonds & Isaac, 2007) et al., 1999; Kennedy, 1999; Geraci & Lounsbury, 2002) † Higher exposure to contaminants stocked in blubber and † Animals like bottlenose dolphins (Tursiops truncatus)in the Gulf of Mexico, living in coastal zones where mobilized during starvation (Aguilar et al., 1999), that alter reproductive, endocrine and immune systems (Fuller & seawater temperature variation is enhanced (IWC, 1997) Hobson, 1986; Aguilar & Borrell, 1994; Ross et al., 2000) † Cold-water species like white-beaked dolphins † (Lagenorhynchus albirostris) in north-western Scotland Stranding of hundreds of grey whales (Eschrichtius robustus) in eastern America (Grebmeier & Dunton, 2000; Moore whose relative abundance and occurrence decreased whereas warm-water common dolphins (Delphinus et al., 2003; Gulland et al., 2005) † delphis) increased in this region (MacLeod et al., 2005) Episodic shifts in cetacean populations’ abundance and distribution in the Gulf of Maine. Concerned species: † Long-finned pilot whale (Globicephala melas) population structure in the North Atlantic and short-finned pilot humpback and fin whales replaced by right and sei whales (Balaenoptera borealis); white-beaked dolphins whales (Globicephala macrorhynchus) distribution in (Lagenorhynchus albirostris) replaced by Atlantic white-sided Japan (Fullard et al., 2000) dolphins (Lagenorhynchus acutus); and harbour porpoise (Phocoena phocoena)(Kenneyet al., 1996; Palka et al., 1997) Effects on prey availability affecting the distribution and † geographical range of: Episodic shifts in the inshore incidence of Pacific white-sided dolphins (Lagenorhynchus obliquidens) in British Columbia † Long-finned pilot whales in the Faeroe Islands (Bjorge, (Morton, 2000) 2002) † Disappearance of common dolphins (Delphinus delphis) † Bottlenose dolphins in north-east Scotland (Wilson et al., along the north-eastern coast of Florida (Caldwell & 2004) Caldwell, 1978) Effects on prey availability affecting the reproductive success of: † North Atlantic right whales (Eubalaena glacialis) (Greene & Pershing, 2004) Precipitation Effects of warmer seawater temperature and nutrient Effects of pollutants inputs into coastal waters due to increased and extreme enrichment due to increased runoff: runoff and flooding events: weather † Increase in the frequency and strength of toxic algal bloom † Impacts on coastal marine mammals and other organisms occurrence episodes triggering lethal poisoning in marine mammals (including prey species) (e.g. MacGarvin & Simmonds, (Herna´ndez et al., 1998; Burns, 2002; Geraci & 1996; Orr et al., 1992; Learmonth et al., 2006) Lounsbury, 2002) † Increase in eutrophication events affecting marine Effects on oceanic properties (coastal waters salinity, circulation, organisms dynamics like phytoplankton in the North etc) affecting: Sea (Edwards et al., 2001) † The distribution and abundance of cetaceans’ prey species (e.g. MacGarvin & Simmonds, 1996; Learmonth et al., 2006) Sea level rise Flooding of coastal habitats and impacts on species that † Grey and humpback whales (IWC, 1997) depend on coastal areas (as breeding, nursing, feeding, mating, resting zones) like: † Mediterranean monk seals (Monachus monachus) breeding in caves or on small beaches (Harwood, 2001; Wu¨rsig et al., 2002) Ocean Changes in current patterns and oceanic fronts directly Changes in water mass formation, mixing and upwelling pattern circulation affecting: affecting: † The distribution of the majority of tropical and temperate † Marine biodiversity and ecosystems (IPCC, 2001a) cetaceans (Worms et al., 2005; Learmonth et al., 2006) like † The distribution, abundance and migration of plankton, fish sperm whales associated with the Antarctic convergence and cephalopods (e.g. Planque & Taylor, 1998; Waluda in the Southern Ocean (Boyd, 2002) et al., 2001; Walther et al., 2002) affecting cetaceans like minke whales (Bjorge, 2002)

Continued 188 d.d. gambaiani et al.

Table 2. Continued.

Global Impacts on marine mammals with some examples of species most likely to be, or, already affected warming- induced changes

Salinity Direct affects on cetaceans epidermis generating: Effects on prey species through changes in oceanic properties or † More important skin lesions in bottlenose dolphins living low salinity tolerance like: in salted and cold waters (Learmonth et al., 2006) † Most cephalopod species (e.g. De Heij & Baayen, 2005) † Animal increased stress and susceptibility to diseases or † Coastal phytoplankton of the Antarctic peninsula (Moline anthropogenic impacts (Wilson et al., 1999) et al., 2004)

Seawater pH Increased CO2 affecting metabolic function, growth and † High metabolic rate species like ommastrephid squids (e.g. reproduction of water-breathing organisms (e.g. Po¨rtner Illex illecebrosus) (Learmonth et al., 2006) et al., 2004; Orr et al., 2005; Royal Society, 2005) Large-scale Increase in frequency of large-scale atmospheric events † Sperm whale (Physeter macrocephalus) reproductive success atmospheric (Timmermann et al., 1999) affecting the distribution, in the Galapagos (Whitehead, 1997) patterns growth, abundance and recruitment of marine organisms † Dusky dolphins (Lagenorhynchus obscurus) reproductive occurrence (IPCC, 2001a; Stenseth et al., 2002) including: success in Peru (Manzanilla, 1989) and strength † Market squid (Loligo opalescens), a short-finned pilot † Short-beaked common dolphins (Delphinus delphis) and (e.g. North whales’ prey (Shane, 1995) other Delphinidea distribution in New Zealand (Gaskin, Atlantic † North Atlantic cod (Gadus morhua) (Stenseth et al., 2002) 1968; Neumann, 2001) Oscillation † North Atlantic copepods Calanus finmarchicus and † Southern right whales breeding success (Leaper et al., 2006) (NAO), El Calanus helgolandicus (Planque & Taylor, 1998; Beare † Californian coastal bottlenose dolphins geographical range Nin˜o, et al., 2002; Beaugrand & Ibanez, 2004) (Wells et al., 1990; Wu¨rsig et al., 2002) Southern † Grey whale high mortality rate and low recruitment in the Oscillation, Large-scale atmospheric events’ impacts on prey availability Pacific Ocean (Le Boeuf et al., 2000) Pacific and thus affecting: † Pacific killer whales (Orcinus orca) and Atlantic bottlenose Decadal † Community structure of short-finned pilot whale dolphins social organization and behaviour (Lusseau et al., Oscillation (Globicephala macrorhynchus) being replaced by Risso’s 2004) (PDO)) dolphins (Grampus griseus) in southern California † North Atlantic right whales breeding success (Simmonds & (Shane, 1995; Wu¨rsig et al., 2002) and in the Gulf of Isaac, 2007) Mexico (Jefferson & Schiro, 1997) † Sperm whale in the north-east Atlantic (Robinson et al., 2005) † Probably, the North Atlantic cetaceans in general, in response to the NAO (Learmonth et al., 2006)

Mediterranean monk seal (Monachus monachus) (Van de alteration of ocean circulation is likely to modify larval trans- Bildt et al., 1999). A worldwide increase in mass mortality port processes, species dispersal and recruitment, and impact events in marine mammals has been reported by Simmonds krill population dynamics (Harley et al., 2006). & Mayer (1997). According to Einarsson (1945), in the north Atlantic, the It is interesting to note that, as cetaceans are long-lived, optimal temperature range of Meganyctiphanes norvegica is slow-reproducing animals (generally producing one offspring between 2 and 158C with a high mortality rate above 158C per female every 2–3 years), when a population is severely (Buchholz et al., 1995). This thermal limit is 188C for the diminished by a virus or other agents, recovery may be slow Mediterranean population (Fowler et al., 1971). Salinity and such species can relatively easily become endangered change is also likely to affect this species which has its toler- (Dhermain et al., 2002; Reeves et al., 2003). ance limit at 20–24 ppm (Forward & Fyhn, 1983). According to Greene & Pershing (2004), in the North Whereas, temperature warming in the north-east Atlantic Atlantic Ocean, the effects of global warming on the abundance has led to the migration of several marine organisms to north- of C. finmarchicus strongly influence right whale (Eubalaena ern latitudes (e.g. Beaugrand et al., 2002), in the glacialis) calving rates. A similar situation is likely to take Mediterranean Sea, Meganyctiphanes norvegica will not be place in the Mediterranean. For instance, Meganyctiphanes able to extend its range northward because of the land norvegica, which constitutes the only known food supply of barrier and is likely to share its environment with more ther- the fin whales (Balaenoptera physalus) in this region, is at the mophilic invasive species in the future. northern limit of its distribution (Besson et al., 1982; Viale, Furthermore, calcifying organisms including some phyto- 1985; Orsi Relini & Giordano, 1992; Orsi Relini et al., 1994; and zooplankton species are likely to be affected by acidifica- Gannier, 1995, 1997; Forcada et al., 1996; Astruc & tion (Royal Society, 2005) and a possible temporal mismatch Beaubrun, 2001; Notarbartolo di Sciara et al., 2003) and thus, may result between Meganyctyphanes norvegica and phyto- in case of unsuitable environmental properties, will not be plankton blooms, its food supply, with protential conse- able to move northward because of the physical land barrier. quences for predators including the endangered bluefin tuna The distribution and abundance of this euphausid are cor- (Thunnus thynnus), albacore tuna (Thunnus alalunga) related with specific hydrobiological parameters (e.g. seawater (Quynh, 1978), squid (Illex coindetii) (Sanchez, 1982) and temperature, salinity, food availability and current patterns) the Mediterranean fin whale population. (Pustelnik, 1976; De la Bigne, 1985; Macquart-Moulin & Mediterranean fin whale distribution can be expected to be Patriti, 1996; Velsch, 1997). Climate change-induced affected by food availability (Littaye et al., 2004) and since environmental threats to mediterranean cetaceans 189

Mediterranean fin whales are genetically and reproductively presenting as skin lesions) and make cetaceans more suscep- isolated from those of the Atlantic (Be´rube´ et al., 1998), they tible to diseases or anthropogenic pressures (Wilson et al., are regarded as more vulnerable to environmental pressures, 1999; Learmonth et al., 2006). These pressures, including inci- including global warming (Dhermain et al., 2002). dental capture in fishing nets, noise and chemical pollution By affecting the distribution and abundance of cetacean could have synergistic or cumulative impacts with climate prey species, climate change is likely to trigger dietary compe- change. tition between species, and could then cause inter- and Finally, the increased frequency of poisonous algal blooms, intra-species competition between Mediterranean cetaceans. like dinoflagellates, which often generate brevetoxins, has been This is particularly true for striped dolphins and common dol- correlated with the collapse of several marine species includ- phins (Aguilar, 2000), whose food web dynamics have been ing cetaceans like striped dolphins in the Mediterranean Sea affected by recent temperature changes (Bearzi et al., 2003). (Geraci et al., 1989; Fitzgerald, 1991; Burns, 1998, 2001, In addition, a decrease of prey species may increase cetacean 2002; Balmer-Hanchey et al., 2003; Danovaro, 2003). Global mortality rates and vulnerability to diseases as a consequence of warming, which is likely to encourage this phenomenon, reduced immune function, as in the Mediterranean striped could indirectly, yet profoundly affect cetaceans. dolphin epizootic outbreak in the 1990s (Aguilar & Raga, 1993; Bearzi, 2002). CONCLUSION climate change combined with fisheries This review illustrates the different linkages existing between pressure climate and biodiversity. It illustrates the urgent necessity The combination of climate-induced impacts with other anthro- for more integrated regulations for the protection of marine pogenic impacts like overfishing is likely to impact cetaceans biodiversity. Similarly, the development of exemplary and (CIESM, 2000; Bearzi, 2002). Several cetacean species such as reproducible projects aiming to reduce greenhouse gas emis- coastal bottlenose dolphins and common dolphins are already sions should be supported. competing with fishermen for prey species exploited by fisheries By influencing seawater properties, climate change can (Bearzi, 2002; Abad et al., in press). As previously observed prey alter ecological interactions between trophic levels and is species distribution and abundance could be severely affected by likely to disrupt overall ecosystem function. Climate change global warming. The diminution of fish stocks is likely to result is affecting and will continue to affect marine ecosystems. in a stronger competition with fishermen and in higher risks of Although recent climatic anomalies represent only a small harassment of dolphins by fishermen (Northridge, 1984; part of the forecasted changes, they have already generated UNEP/IUCN, 1994; Fertl & Leatherwood, 1997; Bearzi, 2002). important responses in marine ecosystems. Marine biodiver- According to Bearzi et al. (2004), global warming is today a sity, including cetaceans, is highly vulnerable to environ- major concern for Mediterranean common dolphin which mental alteration, and can be significantly and irremediably feed on species that are targets of fisheries, such as affected by even small temperature changes. Combined with European anchovy, European pilchard (Sardina pilchardus), other anthropogenic pressures, climate change is likely to round sardinella (Sardinella aurita) and sprat (Sprattus sprat- impact the survival of some rare and endangered marine tus) (Orsi Relini & Relini, 1993; Boutiba & Abdelghani, 1995; flora and fauna, and may also threaten many other species. Birkun, 2002; Bearzi et al., 2003) and that could be affected by For example, some cetacean species that inhabit restricted global warming. For instance, in the Black Sea, the two mass geographical zones, with no option to shift their range, or mortality events involving common dolphins (Delphinus those unable to effectively switch prey species if necessary delphis), in 1990 and 1994, coincided with the decline of may be adversely affected by climate change. As well as European sprat and anchovy stocks, their main prey species impacts on populations of marine biota, the physiology of (Krivokhizhin & Birkun, 1999; Birkun, 2002). The combi- individual organisms may also be severely affected, either nation of several factors, including seawater eutrophication, directly or indirectly, by climate change, for example which is likely to be encouraged with global warming, and through ocean acidification. over-fishing, were responsible for the rapid decline of sprats Today, the International Whaling Commission considers and anchovies (Zaitsey & Mamaev, 1997). global warming as a major issue (IWC, 1997) and is in the process of establishing a special workshop to examine its other possible climate change-induced impacts and as Baker (in Burns, 2001) said: ‘While we impacts on cetaceans debate the limits that should be placed on whaling in order to protect the status of the stocks, a silent menace threatens Climate change is likely to affect cetacean populations in to destroy the populations we strive to protect’. Climate several other ways (Figure 2). 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